Memoirs of the Queensland Museum

VOLUME 14

PART

MEMOIRS

OF THE

QUEENSLAND MUSEUM

BRISBANE

VOLUME 14

PART 2

MEMOIRS

OF THE

QUEENSLAND MUSEUM

ISSUED BY THE AUTHORITY OF THE MINISTER FOR EDUCATION,
THE HON. J. C. A. PIZZEY

CONTENTS

& a

VOLUME 14 PART 2
(Issued 4th July, 1962)

Page

The Application of the Generic Name Macropus Shaw 1790 and of Other
Names Commonly Referred to the Grey Kangaroo J. H. Calaby,

George Mack and W. D. L. Ride .25

A New Species of Thylacoleo and Notes on Some Caudal Vertebrae of

Palorchestes azael . . Alan Bartholomai . 33

Fossil Marsupials and Cainozoic Continental Stratigraphy in Australia: A

Review J. T. Woods .41

THE APPLICATION OF THE GENERIC NAME MACROPUS SHAW
1790 AND OF OTHER NAMES COMMONLY REFERRED TO
THE GREY KANGAROO

J. H. Calaby, Wildlife Survey Section, C.S.I.R.O., Canberra
George Mack, Queensland Museum, Brisbane
W. D. L. Ride, Western Australian Museum, Perth

There has been considerable discussion in recent years as to the identity
of the kangaroos collected in the vicinity of the Endeavour River, north-east
Queensland in 1770 by Captain James Cook’s party. Three specimens were obtained,
weighing respectively 8, 38 and 84 pounds. From an account of this material,
Muller (1776) described the kangaroo ( Mus canguru) mentioning only the 38 pound
specimen. This is the holotype of Mus canguru Muller and it has been generally
accepted that the specimen was a grey kangaroo.

Iredale and Troughton (1925), as a result of examining the description
contained in Solander’s unpublished manuscript (1768-1771), expressed doubt that
the original specimen described and named Mus canguru by Muller (1776) was a grey
kangaroo. They suggested that it might have been a wallaroo of the robustus group.
Later, the same authors (1937) endeavoured to show that it was really a whiptail
or pretty- faced wallaby ( Wallabia elegans). Again, this conclusion was based on
Solander’s manuscript, aided by the purchase of two skins collected near Cooktown
on the Endeavour River, one a whiptail wallaby and the other an antilopine wallaroo.
It is difficult to understand this decision. Solander’s description is a composite one ;
parts of it (one of the weights, some measurements, a sex) can clearly be related to
the holotype, while other parts have been taken from other specimens.

Raven (1939) then discussed the matter and held that the earlier revisers
were correct in indentifying the first described specimen as a grey kangaroo. Morrison-
Scott and Sawyer (1950) provided excellent support for this view, and they produced
additional pertinent evidence that the type of Mus canguru Muller was a young
grey kangaroo. These authors published (1) two outline sketches (indeterminable*)
of an entire kangaroo made by Parkinson, artist on board the Endeavour, (2) wash

* We have examined a coloured transparency (supplied through the Mitchell Library,
Sydney) of a painting of a kangaroo by Stubbs which is probably the original of the figure in
Hawkesworth (see Lysaght 1957). We are unable to identify it specifically with any species
which nowadays occurs at Cooktown. Any attempt to establish that this figure and painting
are of the holotype would be pointless. Muller specifically nominated the holotype by reference
to its weight and there is no evidence that this figure represents the nominated animal.

B

26

MEMOIRS OF THE QUEENSLAND MUSEUM

drawings by Nathaniel Dance of a skull, which can be identified as that of Macropus
robustus, a grey wallaroo ; and (3) a photograph of a skull of a young grey kangaroo
which they designated the “ photo- lectotype ” of Macropus canguru (Muller). This
last skull was collected on Cook’s voyage and was given to John Hunter by Banks,
a member of Cook’s party. It is No. 1732 in Owen’s catalogue (1853) and No. 3703
in Flower’s catalogue (1884). It is the skull of a young grey kangaroo and we believe
that it is from the 38 pound specimen collected near the Endeavour River. Both
Owen and Flower described this skull and its dentition.

Morrison -Scott and Sawyer have suggested that the largest of the three
specimens collected, weighing 84 pounds, was a wallaroo of the Macropus robustus
group, and now represented by the Dance drawings. The presence of a specimen
of the wallaroo in Cook’s collection would account for the statement by Solander in
his composite description that the internarial area of the kangaroo was naked.
A juvenile grey kangaroo from Cook’s voyage (presumably the specimen which weighed
8 pounds) was recorded by Gray (1843) as being preserved in spirits in the British
Museum. It is no longer in the collections. The skull of the third animal, which was
in the collections of the Royal College of Surgeons in London, was destroyed by
a bomb and is now represented by a photograph in the British Museum and
undoubtedly it is the skull of a young grey kangaroo, shown by us here to be from
an animal weighing in the vicinity of 38 pounds, and certainly not 84 pounds.

The instability which now exists in the nomenclature of these well known
kangaroos and wallabies is undesirable and can best be removed by proving the
specific identity of the holotype which was nominated by Muller. Clearly, no
consideration need be given to the 8 pound juvenile grey kangaroo which was in the
British Museum ; the holotype is either the Hunterian specimen or another, probably
represented by the specimen drawn by Dance. It must be emphasised that Solander’s
composite description which was not published until 1925 is of no nomenclatural
importance.

TOPOTYPICAL MATERIAL

Morrison-Scott and Sawyer lacked material from the type locality, and with
this in mind, a party from the Queensland Museum, consisting of D. P. Vernon,
S. Breeden and M. E. McAnna, collected in the vicinity of Cooktown, Endeavour
River, during October and November, 1960. Among the specimens obtained are
five grey kangaroos, seven grey wallaroos, six antilopine wallaroos, seven whiptail
wallabies and ten agile wallabies.

A young male grey kangaroo (J.10749), collected at Kings Plains, November
24, 1960, by Vernon and Breeden, is nearest in weight to the 38 pound animal of
the Endeavour party. Although it weighed 55 pounds, including viscera and stomach
contents, its dental age is precisely that of the 38 pound animal. Another male

APPLICATION OF GENERIC NAME MACROPUS SHAW 1790

27

(J. 10750), weighing 90 pounds, including viscera and stomach contents, was collected
at the same locality. This specimen has all molars fully erupted, and not only has
dP 4 been shed, but its successor (P 4 ) also has been shed on the right side and is close
to being shed on the left. It will be clear from these remarks that the 84 pound
Endeavour specimen cannot have been the skull given by Banks to Hunter, and on
the evidence afforded by the specimen (J. 10749) in the Queensland Museum, it must
have been the 38 pound animal. Furthermore, in addition to agreeing exactly in
dental age, the limb measurements of the young male (J. 10749) alone agree closely
with those provided by Muller when describing Mus canguru.

The exact stage of dental eruption reached by the Hunterian (38 pound)
specimen was well described by both Owen and Flower. P 3 has been shed, dP 4 was
about to be replaced by P 4 ; M 1 , M 2 and M 3 were in place in the maxilla and M 4 was
still in its crypt. The dentition of J. 10749 is the same as will be seen from the
accompanying plates.

It was not possible to establish on the recent Cooktown material the range
of variation in weight which can be expected at this stage of dentition. However,
for this purpose, Mr. W. H. Butler, Associate of the Western Australian Museum,
collected a series of male grey kangaroos from a single population at Congelin,
south-west Australia. Three of these were at the same dental age as both of the
above, and their weights were 63, 68 and 79 pounds, a weight range of 16 pounds.
The south-west grey kangaroo is a heavier animal than the north-east Queensland
form, but the difference in minimum and maximum weights is similar to the difference
between the 38 pound specimen of Cook’s party and the Queensland Museum
example, J. 10749. It would appear that the 17 pounds difference is not significant.

Finally, in order to satisfy ourselves as to the probability of the identification
of the skull of the Dance drawing with the 84 pound specimen, this (as reproduced
in Morrison -Scott and Sawyer) has been carefully compared with skulls from the
Endeavour River of the whiptail wallaby, the agile wallaby, the antilopine wallaroo,
the grey wallaroo and the grey kangaroo, and it is clearly a specimen of the grey
wallaroo. At first sight, the third upper incisor appears unusual until it is realised that
Dance, in order to show the presence of the faint groo\e in the outer surface of the
posterior lobe, has slightly overemphasized it. The skull drawn by Dance has a
sectorial tooth followed by three fully erupted molariform teeth, and a partly open
alveolus is shown behind the last molariform tooth. The cementum area of the root
of the third incisor is well exposed and the pronounced supraorbital crests become
confluent above the temporal fossa to form a sagittal crest. These growth characters
support our identification of the teeth as P 4 , M 1 , M 2 , and M 3 (with M 4 in its crypt).
Hawkesworth says of the 84 pound animal, that it was “ not at its full growth, the
innermost grinders not yet being formed.” A male specimen of the grey wallaroo
at this dental age (J. 10738) was collected at Annan River, 17 miles south of Cooktown
and it weighed 70 pounds.

28

MEMOIRS OF THE QUEENSLAND MUSEUM

The skull of the wallaroo drawn by Dance is very probably that of the
84 pound animal shot by Lieutenant Gore on July 27th, 1770. Its presence in the
series would explain the presence of the characters which are atypical of the grey
kangaroo in the composite description of Solander (e.g. the naked internarial region).
The grey wallaroo is the only <c grey ” macropod in the area which possesses these
characters and at the same time achieves a weight of 84 pounds. Further it should
be noted that, of the five species collected at the Endeavour River in 1960, the
male grey wallaroo is the only male form which is sufficiently nondescript externally
to be included with a series of grey kangaroos without remark. Cook’s party obtained
two grey kangaroos, and male specimens of species other than the grey wallaroo are
either widely different from the grey in colour or are, even to the untrained eye,
brightly and obviously ornamented.

WHIPTAIL WALLABY

It has already been stated that the final decision of Iredale and Troughton
that Mus canguru Muller was based on a whiptail wallaby was not derived from an
examination of Muller’s description, but from an examination of a description
contained in a manuscript by Solander, now in the British Museum. This description
included both the male and female genitalia and the pouch, and the weights of
three animals were given. The main characters stated by Solander and used by
Iredale and Troughton were that the area between the nares was naked and that
the third incisor was bilobed, broad from side to side and with smaller anterior
lobes. As Morrison-Scott and Sawyer have pointed out, Solander’s description
of the incisors accords neither with the grey kangaroo nor the whiptail wallaby,
but it is in accord with the condition in the wallaroo.

Solander also stated “ Par intimum Molarium diu intra alveolus sous latit,
in junioribus non discernandum,” implying that both the 38 and 84 pound specimens
had unerupted last molars. He referred to the 38 pound example as a male of two
or three years, and to the 84 pound animal as “ adultus.” Similarly, Hawkesworth
described the 38 pound animal as “ a young one, much under its full growth. ”
Elsewhere in the same work Hawkesworth made it clear that the term “ full growth ”
referred to the possession of fully erupted molar teeth since he said of the 84 pound
animal “ We found that this animal was not at its full growth, the innermost
grinders not being yet formed.”

All male whiptail wallabies between 30 and 50 pounds in weight collected
on the recent field trip to the Endeavour River had erupted last molars, as was to be
expected. Externally, each individual has a prominent white stripe, approximately

APPLICATION OF GENERIC NAME MACROPUS SHAW 1790

29

12 mm. wide, extending from the nares to behind the eye on each side of the face ;
another prominent white stripe, crescent-shaped, on each thigh ; and the fur under
the head, centre of throat and entire abdomen is white. It is most unlikely that those
who examined and described the first collected Macropodidae from Australia would
ignore these vivid markings if the whiptail wallaby was represented in the collection.

CONCLUSION

From the above evidence, together with the evidence provided by Morrison-
Scott and Sawyer, it would appear that the following specimens were collected in the
vicinity of the Endeavour River, north-east Queensland, in July, 1770 by Cook’s
party.

(1) A grey wallaroo, Macropus robustus, weighing 84 pounds. It is likely

that from this specimen Solander in manuscript described the area
between the nares as naked. Now apparently represented by a wash
drawing of the skull by Nathaniel Dance in the British Museum.

(2) A young grey kangaroo, weighing 38 pounds. This specimen, which

had P 4 and M 4 still in their crypts, was described by Hawkesworth
(1773) and described and named Mus canguru by Muller in 1776.
Now represented by a photograph of the skull in the British Museum.

(3) A juvenile specimen, weighing 8 pounds, a grey kangaroo. Recorded

as preserved in the British Museum (Gray, 1843), but not now in the
collections.

CONSERVING THE GENERIC NAME MACROPUS

Macropus is currently the generic name of the large-sized kangaroos, including
the grey kangaroo, and it has been applied in this way for more than one hundred
years. The genotype is Yerboa gigantea Zimmermann (1777) which is accepted as
a synonym of Mus canguru Muller (1776).

The view of Iredale and Troughton is that the type of Mus canguru Muller
was a whiptail wallaby, at present generally known as Wallabia elegans. To those
who accept this view, the generic name Macropus must be used for the large-sized
wallabies, and another name will have to be found for the large-sized kangaroos.

30

MEMOIRS OF THE QUEENSLAND MUSEUM

Since the case presented here rests, to some extent, on probability, in order
to stabilize both the generic and specific names involved, we have decided to approach
the International Commission for Zoological Nomenclature with the following
request : —

(1) That the specimen, the skull of which was given by Sir Joseph Banks
to John Hunter and became No. 3703 in the collections of the Royal
College of Surgeons, London, be declared the holotype of Mus canguru
Muller 1776 ; that since this specimen has been destroyed, it be
replaced by a neotype, a grey kangaroo, Queensland Museum
No. J.10749, male, skin and skull, collected at Kings Plains, 20 miles
south of the Endeavour River, November 24, 1960, by D. P. Vernon
and S. Breeden.

(2) That a procedure be adopted to make the name Yerboa gigantea
Zimmermann 1777 an objective synonym of Mus canguru in accordance
with the current usage of these two names.

(3) That the name Macropus major Shaw 1800 be conserved for the grey
kangaroo with type locality Sydney as restricted by Iredale and
Troughton (1934). According to Article 7 2d of the International Code,
Macropus major has the same type specimen as Yerboa gigantea and
the Plenary Powers will have to be exercised in order to validate the
action of Iredale and Troughton.

If the Commission accepts this recommendation, the generic name Macropus
will continue to be available for the large-sized kangaroos ; M . canguru will be the
name of the grey kangaroo ; and major will be available for the grey kangaroo of
Sydney and beyond should it prove to be subspecifically distinct from the Endeavour
River, north-east Queensland grey kangaroo.

It is a pleasure to acknowledge the assistance rendered by the staff of the
Mammal Department, British Museum (Natural History), in particular Miss Jean
Ingles and Dr. Gordon Corbet ; the Fisheries and Wildlife Department, Victoria,
especially Mr. John McNally ; Mr. E. H. M. Ealey of the Department of Zoology
and Comparative Physiology, Monash University ; and Mr. W. H. Butler, Associate
of the Western Australian Museum.

APPLICATION OF GENERIC NAME MACROPUS SHAW 1790

31

LITERATURE CITED

"Flower, W. H., 1884. Catalogue of specimens ... in the Museum of the Royal College
of Surgeons of England, 2, p. 708.

Gray, J. E., 1843. List of . . Mammalia in the British Museum, p. 87.

Hawkesworth, J., 1773. An account of the Voyages undertaken . . . for making Discoveries

in the Southern Hemisphere ... by Captain Cook, 3, p. 577 (1st Ed.), p. 173
(2nd Ed.). London.

Iredale, T., and Troughton, E.LeG., 1925. Captain Cook’s Kangaroo. Av&t. Zool., 3, p. 311.

, 1934. A Checklist of the Mammals Recorded from Australia, p. 54.

, 1937. The identity of Cook’s Kangaroo. Rec. Aust. Mus,, 20, p. 67.

Lysaght, A., 1957. A clue to the mystery of Captain Cook’s Kangaroo. The New Scientist, 3, p. 17.

Morrison- Scott, T. C. S., and Sawyer, F. C., 1950. The identity of Captain Cook’s kangaroo.
Bull. Brit. Mus. (Nat. Hist.), Zocl., 1, p. 43.

Muller, P. L. S., 1776. Syst. Nat. (Linne), Suppl., p. 62.

Owen, R., 1853. Descriptive Catalogue of the Osteological Series . . . Museum

Royal College of Surgeons of England, 1, p. 322. London.

Raven, H. C., 1939. The identity of Captain Cook’s Kangaroo, J. Mammal ., 20, p. 50.

Shaw, G., 1800. Gon. Zool., 1, pt. 2, p. 505.

-Zimmermann, E. A. W., 1777. Specimen zoologiae geographicae, Quadrupedum domicilia et
migrationes sisteus & etc., p. 526. Leyden.

EXPLANATION OF PLATES
Macropus canguru. Grey Kangaroo
Plate V. Doral view of skull.

Plate VI. Ventral view of skull.
Plate VII. Lateral view of skull.
Plate VIII. Mandible.

All views of same skull, J. 10749, male (Q.M.). All natural size
Specimen from Kings Plains, 20 miles south of Cooktown, N.E. Queensland.

MEMOIRS OF THE QUEENSLAND MUSEUM, Vol. XIV, Part II, Plate V

mm m

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MEMOIRS OF THE QUEENSLAND MUSEUM, Vol. XIV, Part II, Plate VI

MEMOIRS OF THE QUEENSLAND MUSEUM , Vol. XIV, Part II, Plate VII

Ouecmlrtod Museum

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MEMOIRS OF THE QUEENSLAND MUSEUM , Vol. XIV, Part II, Plate VIII

33 >

A NEW SPECIES OF THYLACOLEO AND NOTES ON SOME
CAUDAL VERTEBRAE OF PALORCHESTES AZAEL

Alan Bartholomai
Queensland Museum

The Darling Downs area of south-eastern Queensland consists mainly of
extensive fluviatile deposits of Upper Cainozoic Age. Woods (1960) indicated that
the Chinchilla Sand, developed in the valley of the Condamine River in the north-
western Darling Downs, was probably deposited earlier than the Pleistocene alluvia
to the south-east.

Specimens of the genus Thylacoleo Owen in the collections of the Queensland
Museum were revised by Woods (1956) and those from the south-eastern Darling
Downs were attributed to T. carnifex Owen. A small proportion of fragmentary
material from the Chinchilla district was tentatively separated as possessing features
sufficiently distinct to constitute a possible new species. Recently, a partial right
mandibular ramus was collected from the Chinchilla Sand at the Chinchilla Riflo
Range (Rifle Range Number 78, parish of Chinchilla), supporting the view that the
variations are characteristic, and the material is here described as a new species.

All measurements are in millimetres.

THYLACOLEO CRASSIDENTATUS sp. nov.

(Figures 1, 2)

Material. F.3565, holotype. A partial right mandibular ramus with l t
broken, P 3 - M 2 , young adult. Chinchilla Sand, at 363677 Chinchilla 4 mile military
map, possibly Pliocene.

Specimens from the Chinchilla Sand at Chinchilla, north-western Darling
Downs : F.2957, partial right mandibular ramus, no teeth preserved. F.296R

partial left mandibular ramus with both P 3 and M x broken, very aged. F.2962,
partial right mandibular ramus with P 3 , adult. F.2960, partial left mandibular ramus
with M 2 , aged. F.2964, partial left mandibular ramus with l ± broken, P 3 , juvenile.
F.2963, broken left P 3 , adult. F.2495, partial left mandibular ramus with P 3 , adult.
F. 2941, partial right maxilla with both P 3 and M 1 broken, adult. F.2954, partial
right maxilla with both P 3 and M 1 broken, adult. F.2955, partial right maxilla with P 3 ,.
adult.

34

MEMOIRS OF THE QUEENSLAND MUSEUM

Specimens from the Darling Downs : F.3569, partial right mandibular ramus
with both 1 1 and P 3 broken, M 1? adult. F.3570, partial left mandibular ramus with
Mj, adult. F.3571, partial left mandibular ramus. F.3572, partial left mandibular
ramus with both P ;j and M x broken, aged.

Measurements of Mandibles

Specimen

Length of
crown of

Breadth of P 3
above

posterior root

Length of
crown of Mi

Breadth below
metaconid
of M t

Angle between
1 1 and base
of mandible

Holotype, F.3565

370

141

15-5

11-3

42°

F.2962

35-6

13-6

—

—

—

F.2964

36-2

13-5

—

—

—

F.2495

41-4

14-3

—

__

—

F.2961

—

—

—

—

39°

F.3569

—

—

13-3

10-0

42°

F.3570

—

—

14-9

10-8

—

F.3572

—

—

15-5

11-5

—

Ramus deep, strong, particularly robust in region of implantation of cheek teeth ;
longitudinal axis very slightly convex laterally ; symphysis unfused ; symphysial plane short,
deep, subequilaterally triangular, somewhat upturned. Fossa subalveolaris deep, confluent ;
mental foramen prominent, ventral to anterior margin of P 3 and antero-dorsal to junction of
-anterior margin and inferior surface of ramus at blunt angle ; lateral alveolar walls of P 3 thinning,
roots becoming exposed with age, with simultaneous development of inter-rootial depression.
Ramus ascending at low angle posterior to smooth, weak diagastric process ventral to M 2 ;
postalveolar ridge not prominent, ascending gradually posteriorly, disappearing on mesial wall
of large coronoid process. Process diverging from line of ramus, directed antero-posteriorly ;
antero-dorsal margin ascending at approximately 45°, grooved, laterally flanged ; flange continuing
ventrally on body of ramus limiting large, anteriorly deep, ectocoronoid fossa. Wall of fossa
perforated by masseteric foramen opening into inferior dental canal close to mandibular foramen.
Condyle, postero-mesial angle of ramus and posterior portion of coronoid process not preserved.

Lower median incisor broken, but enough remains to indicate the presence of shallow,
posterior longitudinal furrows ; lingual furrow best developed. 1 x making an angle of approxi-
mately 42° with base of mandible. P x and P 2 not retained ; alveoli small, mesiad to anterior
root P 3 . Third premolar elongate, sectorial, very convex laterally, deeply rooted ; roots directed
somewhat dorso-laterally ; base of crown swollen ; enamel finely ridged vertically, particularly
•on lingual surface, thickened to flange on anterior edge ; crown broadest above posterior root,
not tapered posteriorly, asymmetric, with labial face set at much higher angle than lingual ;
prominent antero-lingual buttress associated with anterior cuspid. Main surface of wear near
planar, developed ventro- lateral to cutting edge. Enamel also removed labially at base of crown.
Molar series reduced. Mj relatively large, subtriangular ; anteriorly with metaconid high, flanged,
with short longitudinal cutting edge in functional continuity with that of P 3 ; base of crown broad
-anteriorly due to development of prominent labial buttress ; crown posteriorly reduced with low
median ridged area and shallow, finely ridged, dorso- labial fossette. Facet of wear developed

A NEW SPECIES OF THYLACOLEO AND PALORCHESTES AZAEL

35-

in continuation with that of P 3 , and another directed postero-laterally ; birooted, anterior root:
much larger than posterior. M a small, with single root partially divided lingually, not functional;,
crown with shallow, faintly ridged dorsal depression.

Figure 1. Thylacoleo crassidentatus sp. nov. F.3565, holotype. Right ramus, lateral and occlusal?
views. Natural size.

Woods (1956) compared the fragmentary Thylacoleo material from Chinchilla
with T. carnifex from the south-eastern Darling Downs and discussed the differences
exhibited. Although only slight dissimilarities are observed in the maxillary remains,,
the mandibles differ considerably.

The most significant feature is the structure of the cheek teeth and, in-,
particular, that of the lower sectorial premolar, where the tooth is posteriorly much
broader and the longitudinal convexity much stronger than in the corresponding P 3 .
of T. carnifex. M x exhibits a difference in relative proportions ; it is comparatively
stout owing to the greater width above both the anterior and posterior roots. The
posterior portion of the crown is comparable with the size of the posterior root and.

36

MEMOIRS OF THE QUEENSLAND MUSEUM

-exhibits a much wider median ridged area together with the development of a shallow,
postero- labial fossette. The second molar, although variable developed, is stronger
than in T. carnifex and its alveolus is partially divided by a vertical ridge on the
lateral wall. In one specimen (F.2960), a depression immediately posterior to the
relatively large second molar is interpreted as the alveolus of a very small third molar.

In addition, the ramus is much wider in the region of implantation of the cheek
deeth and displays a less pronounced longitudinal concavity of the mesial side, while
.a more recumbent lower median incisor is indicated by the reduction of the angle
between the incisor and the base of the mandible.

Figure 2. Thylacoleo crassidentatus sp. nov. Lateral and occlusal views of maxillary fragment
(F.2954). Natural size.

A NEW SPECIES OF THYLACOLEO AND PALORCHESTES AZAEL

37

Owing to the fragmentary nature of the preservation of the maxillary material
in T. crassidentatus relatively few comparative measurements are possible. The third
premolars exhibit no features which differ from those of T. carnifex. The base of the
crown of the first molar in F.2954 is wider than in T. carnifex, while in F.2955 the
tooth is more distinctly tri-rooted, the anterior root being largest, while the posterior
roots are subequal. In addition, F.2954 exhibits a depression immediately posterior
to M 1 and this is considered to represent the alveolus of a small second molar.

It is apparent that the molar series of T. crassidentatus are much less reduced
both in size and numbers than in T. carnifex, a fact in keeping with the suggested time
relationships of the two faunas.

VERTEBRAE OF PALORCHESTES AZAEL OWEN
(Figures 3, 4)

Six caudal vertebrae (F.3564), were recently located in position in the alluvia
of “ Strathmore ” station near Collinsville, north-eastern Queensland, and were
presented to the Queensland Museum by Mr. E. Cunningham. Associated with the
series is a large number of post-cranial fragments together with an incomplete right
mandibular ramus, the lower median incisors, and several maxillary fragments of
JPalorchestes azael. In view of the field association there can be little doubt that the
vertebrae, interpreted as representing the first six of the caudal series, belong to
this species.

Neural arches and zygopophyses are not preserved in first, fourth and fifth
vertebrae, while transverse processes are broken in vertebrae five and six, and are
poorly represented in the first. No chevrons are preserved.

Description

Vertebrae large, stout. All centra short antero-posteriorly, broader posteriorly than long
and broadest in second. Length decreases from first to the fifth, then increases. Centra broadly
convex interiorly ; posterior depth greatest in first caudal vertebra, decreasing regularly to sixth.
Epiphyses firmly fused. Floors of neural canals in anterior vertebrae subdivided by median
ridges, on either side of which foramina pierce centra ; distal centra pierced by one foramen, with
median ridges discontinuous. Neural arches moderately high and broad with neural spine quite
high in second caudal, decreasing in development posteriorly and barely visible as low ridge in
■sixth vertebra. Zygopophyses functional in proximal vertebrae but with anterior zygopophyses
reduced and functionless by sixth vertebra and with posterior zygopophyses lost. Transverse

38

MEMOIRS OF THE QUEENSLAND MUSEUM

processes broadly expanded antero-posteriorly in anterior vertebrae, particularly at extremities,
but are rounded at tips ; processes decrease in transverse diameter and in antero-posterior
expansion to sixth caudal.

Figure 3. Palorchestes azael Owen. Dorsal view of caudal vertebrae (F.3564). One third natural
size.

A NEW SPECIES OF THYLACOLEO AND PALORCHESTES AZAEL

39

Measurements

Caudal Vertebra

Length of
centrum

Posterior
breadth of
centrum

Posterior
depth of
centrum

Breadth across
transverse
process

1

79

86

62

2

71

87

60

272

3

68

80

58

247

4

66

74

56

236

(estimated)

5

66

70

54

—

6

68

69

51

—

In his revision of the species of Palorchestes Owen from south-eastern Queens-
land, Woods (1958) concluded that the genus belongs within the family Diprotodontidae
and not within the Macropodidae as indicated by Owen and later workers. The
erroneous idea of the systematic position of Palorchestes led Owen to attribute a number
of post-cranial macropodid remains to the genus. Fletcher (1945) mentioned additional
post-cranial fossils referred to P. azael in the collections of the Australian Museum,
Sydney, but Woods doubted the identification because of the absence of field
association. Woods has also indicated that the lower incisor found associated with
post-cranial remains from the dune sandstone at Fowler’s Cove, Nepean Peninsula,
Victoria and tentatively referred by Gregory (1902) to this species, is of characteristic
macropodid aspect and referable to one of the large extinct species of Protemnodon
Owen.

Relatively few of the Australian fossil diprotodontids have their caudal
vertebral series sufficiently well known to afford comparisons with those of P. azael.
The posterior breadths of the proximal centra in P. azael and Diprotodon optatus Owen
exceed the lengths, but while the centra of D. optatus are shallow posteriorly owing
to the flattening of the inferior surfaces, those of P. azael are similar in structure to
the macropodids. Gill and Banks (1956) and Scott (1915), show that the centra of
Nototherium tasmanicum Scott are structurally similar to those of D. optatus. The
transverse expansion of the transverse processes and the rounding of the process
extremities in P. azael is similar to that of the other diprotodontids, and is especially
similar to that of N. tasmanicum. The anterior neural canals, however, are comparable
with those observed in the Macropodidae, being high and comparatively broad, but
contrasting markedly with the low, very broad neural canals in D. optatus. In
conjunction with this feature, the zygopophyses in the proximal caudal vertebrae
of D. optatus are much reduced and functionless, while the neural spines are
represented by extremely low tubercles. This is similar to the structure observed
in N. tasmanicum but is in direct contrast to the well-developed, functional
zygopophyses and moderately high neural spines in P. azael.

40

MEMOIRS OF THE QUEENSLAND MUSEUM

Figure 4. Palorchestes azael Owen. Anterior view of second caudal vertebra (F.3564). One third
natural size.

Tedford (1959) recorded a palorchestine diprotodontid from the Etadunna
Formation of possible Oligocene Age (Stirton et al., 1961), at Lake Ngapakaldi and
Lake Kanunka North, Tirari Desert, north-eastern South Australia, and noted that
the tail was long and heavy.

The palorchestine diprotodontids were apparently lightly built, more mobile
grazing animals than other members of the Diprotodontidae. Although the tail of
P. azael differs considerably from that of D. optatus, it does appear to be structurally
closer to that of N. tasmanicum. The structural similarities to the macropodid tail
are believed to be related solely to body form and are not of general systematic
importance.

LITERATURE CITED

Fletcher, H. O., 1945. Palorchestes — Australia’s Extinct Giant Kangaroo. Aust. Mus. Mag., 8,
pp. 361-365.

Gill, E. D., and Banks, M. R., 1956. Cainozoic History of Mowbray Swamp and Other Areas
of North-western Tasmania. Rec. Q. Viet. Mus., 6 (n.s.), pp. 1-42.

Gregory, J. W., 1902. Some remains of an extinct kangaroo in the Dune-Rock of the Sorrento-
Peninsula, Victoria. Proc. Roy. Soc. Viet., 14 (n.s.), pp. 139-144.

Scott, H. H., 1915. A Monograph of Nototherium tasmanicum. Rec. Oeol. Surv. Tasm., 4,pp. 1-47.

Stirton, R. A., Tedford, R. H., and Miller, A. H., 1961. Cenozoic Stratigraphy and Vertebrate-
Paleontology of the Tirari Desert, South Australia. .Rec. S. Aust. Mus., 14, pp. 19-61.

Tedford, R. H., 1959. Notes. Soc. Ver . Paleont. News Bull., No. 55, pp. 23-26.

Woods, J. T., 1956. The Skull of Thylacoleo carnifex. Mem. Qld. Mus., 13, pp. 125-140.

, 1958. The Extinct Marsupial Genus Palorchestes Owen. Mem. Qld. Mus., 13,

pp. 177-193.

— , 1960. Fossiliferous Fluviatile and Cave Deposits. Jour. Geol. Soc. Aust., 7,

pp. 393-403.

41

FOSSIL MARSUPIALS AND CAINOZOIC CONTINENTAL
STRATIGRAPHY IN AUSTRALIA: A REVIEW

J. T. Woods

Geological Survey of Queensland, Brisbane

The geological history of the mammals in other parts of the world suggests
that marsupials entered the Australian Region and became isolated at an early date,
at least by the beginning of the Cainozoic era. However, prior to the discoveries of
the last decade, the early palaeontological record of Australian marsupials was most
unimpressive, with only one species, Wynyardia bassiana, known from lower Tertiary
deposits. Their abundance in Quaternary sediments stood in remarkable contrast.

Since then, the work of Professor R. A. Stirton and his colleagues in the
eastern part of the Lake Eyre Basin in South Australia has provided a major
contribution to the palaeontological record of the marsupials and other vertebrates
in Tertiary time. Other significant discoveries have been reported from the Tertiary
of Victoria, where the association of some of the fossils with marine beds has enabled
more precise dating of the remains. While the bulk of our knowledge is still confined
to upper Cainozoic forms, their potential value in continental stratigraphy is
apparent. At the same time, the variety of these upper Cainozoic marsupials points
to the diversification of the groups in this continent at an early date, and emphasizes
that the paucity of the early Tertiary record will have to be overcome before the
basic phylogeny of these groups can be established.

TERTIARY

The oldest Tertiary marsupial, the age of which can be stated with any
precision, is still Wynyardia bassiana from marine sediments at Fossil Blulf near
Wynyard, northern Tasmania. The skeleton was originally found in a fallen block
of limestone from the “ Turritella Bed,” and the results of the application of the
fluorine test, published by Gill (1957), appear to establish that this bed was its true
provenance. On the basis of its correlation with the Janjukian Stage in Victoria,
an Oligocene age is now generally accepted for the marine section at Fossil Blulf.
Following his re-examination of the skeleton, Wood Jones (1931) concluded that the
affinities of Wynyardia were with the Phalangeridae.

Several fragmentary marsupial fossils have been recovered from shallow
water marine beds and from horizons interbedded with marine sediments in Victoria.
As our knowledge of such forms increases, their occurrence will provide a basis for
correlation of the continental Tertiaries with the marine succession. The oldest of

42

MEMOIRS OF THE QUEENSLAND MUSEUM

these remains is portion of a macropodid femur reported by Glaessner, McGowran,.
and Wade (1960) from sand of Balcombian age (Middle Miocene) above the Bochara
Limestone in Grange Burn, near Hamilton, Victoria. Younger marine sediments in
the same area, the Grange Burn Coquina of lower Pliocene age, have yielded a
fragment of a macropodid mandible, referred by Stirton (1957b) to the subfamily
Sthenurinae.

Three diprotodontid fossils originally found on the beach at Beaumaris,
Victoria, and described by Stirton (1957b) were shown by Gill (1957) to have a
fluorine index comparable with a provenance in the Black Rock Member of the
Sandringham Sand. These marsupials then belong to the Cheltenham “ Stage ” of
upper Miocene age.

Details of the stratigraphic succession and additional information on the fauna
of the Cainozoic sediments of the Tirari Desert in the eastern part of the Lake Eyre
Basin, South Australia, have been recently published by Stirton, Tedford, and
Miller (1961). While the stratigraphic relationships of the various units have been
established through superposition, the authors have expressed difficulty in assigning;
them ages in terms of the conventional epochs of the Tertiary. The oldest unit
recognised, the lacustrine Etadunna Formation, is tentatively assigned to the
Oligocene on the basis of comparative evolutionary studies of the macropodid fauna.
The formation postdates the development of duricrust on representatives of the
Cretaceous Winton Formation and remnants of early Cainozoic fluviatile deposits.

It must be acknowledged that lateritic processes were operative not only over
wide areas in Australia, but at more than one time during the Cainozoic. However,.
Twidale (1956) has postulated that in north-west Queensland uplift and dissection
of a widespread lateritized surface, developed on Cretaceous and probable early
Tertiary rocks, took place approximately in Miocene time. The surface was apparently
of considerable extent in inland Australia, and constitutes the Australian Pediplain
of King (1950), the erosion of which, he claims, was initiated by earth movements in
late Oligocene or Miocene time. It may well be that a tentative Miocene age for
the Etadunna Formation would be more appropriate than a tentative Oligocene age.

The marsupials of the Etadunna Formation recorded by Stirton et al. (1961)<
include a dasyurid, a phascolarctid, Perikoala, previously described by Stirton
(1957a), two macropodids, and a small diprotodontid with possible palorchestine-
affinities.

Lithologically the formation is calcareous in part, with some massive limestones
carrying chert nodules. Paten (1961) has recently discussed the Cainozoic freshwater
limestones and associated sediments, which are widespread in western Queensland,,
but which show their greatest development in the valleys of the Georgina and Burke
Rivers and near Birdsville, all within the present Lake Eyre drainage basin. Their

FOSSIL MARSUPIALS AND CA1NOZOIC CONTINENTAL STRATIGRAPHY 43

section usually includes detritus from older lateritized profiles at the base, and silicifica-
tion frequently occurs towards the surface. Paten has suggested a late Tertiary or
early Quaternary age for these deposits. Vertebrate fossils have been recovered from
only one locality, in the Carl Creek Limestone, near Riversleigh in the Gregory River
valley. This is away from the main occurrences, and the fragmental bones, so far
recovered, are unsatisfactory for close study.

It is possible that deposition of calcareous lacustrine sediments took place
at different times in the Cainozoic over this whole belt. No correlation of any of these
deposits with the Etadunna Formation can be suggested on the available evidence ;
there is no detailed lithological resemblance, but the similarity in gross stratigraphic
relationships indicates that the possibility might be considered in future work.

In the Tirari Desert the next unit recognised by Stirton et al. (1961), the
Mampuwordu Sands, consist of fluviatile sediments deposited disconformably or
possibly unconformably on representatives of the Etadunna Formation. Elements
of the fauna of the stream channel deposits, tentatively placed as lower Pliocene in
age, were described by Stirton (1955). In the recent work the marsupials of the
faunal list now comprise the peramelid Ischnodon, four macropodids including
Prionotemnus , and two diprotodontids, Meniscolophus and a form with affinities to
Euowenia.

Unfossiliferous sandy and argillaceous sediments of the Tirari Formation,
which overlie the Etadunna Formation unconformably, are also tentatively referred
by these authors to the Pliocene. Fossiliferous Pleistocene sediments, the Katipiri
Sands, and younger Quaternary or Recent fluviatile and aeolian sediments complete
the succession in the Tirari Desert, the most complete recognised in continental
deposits of the Cainozoic of Australia.

To date, elements of the fauna of the Mampuwordu Sands have not been
found elsewhere in Australia. Stirton (1955) suggested that Nototherium watutense,
originally described by Anderson (1937) from the Watut River, New Guinea, was
probably referable to Meniscolophus. Information from Dow (1961, personal
communication) indicates that the New Guinea species occurs in sediments of
Pleistocene age, but some occurrences may be older.

Loosely compacted fluviatile and lacustrine sediments which are tentatively
referred to the Pliocene are widespread in Queensland. They comprise the Glendower
Formation and its lithological equivalents, which are mostly unnamed and unmapped,
although the Lynd Formation is known to be extensive in the plains east of the
southern part of the Gulf of Carpentaria. Conglomerates containing pebbles of the
silicified duricrust, commonly known as “ billy,” are usually prominent in the
section, and the sediments appear to have been derived from the dissection of the
extensively lateritized, peneplaned (or pediplaned) middle Cainozoic surface. They,

44

MEMOIRS OF THE QUEENSLAND MUSEUM

in turn, often display broad ferruginous mottling, usually interpreted as a weak
lateritic effect. There is evidence of broad warping, and in places considerable
dissection has followed uplift.

Unfortunately these widespread deposits are usually unfossiliferous, but the
prevalence of leached non-calcareous elastics lessens the chances of preservation of
vertebrate remains. An exception is the Chinchilla Sand (Woods, 1960) which has
been traced on the basis of both lithology and fossil content for a distance of nearly
40 miles in its surface and subsurface extent between Warra and Nan gram Lagoon
in the north-west of the Darling Downs. The maximum thickness is about 100 feet
and calcareous horizons occur. Tortoise and crocodile remains are abundant as well
as marsupials in the extensive vertebrate fauna. There is a striking difference
between the diprotondontid elements of this fauna and those of the superficial
deposits of the eastern Downs. Of these Euryzygoma is absent from the presumably
younger alluvia, while Euowenia and Palorchestes are represented by distinct species.
However superposition has yet to be established.

A few vertebrate fossils are known from deep alluvia, beneath basalts, in
southern Australia. Some of these occurrences are probably upper Tertiary, such
as those at Buninyong, Victoria, at a depth of 238 feet (Gill, 1957), and the Canadian
Lead at Gulgong, New South Wales, at a depth of 130 feet (Dun, 1895). Freshwater
sediments are known to occur below the upper Cainozoic volcanics of north Queensland,
but only plant remains have been recovered from them.

Additional localities listed by Gill (1957) for the possible occurrence of Tertiary
marsupials in southern Australia include One Tree Point, Hobart, and the Geilston
Travertine in Tasmania, the site at Smeaton, Victoria, whence came the dasyurid,
Glaucodon ballaratensis Stirton, and the lacustrine deposits at Coimadai, Victoria.

QUATERNARY

Vertebrate remains assigned to the Quaternary are widespread in fiuviatile,
lacustrine, and cave deposits, while there are more restricted occurrences in spring
deposits, aeolianites, and tuffs. Detailed stratigraphic studies have yet to be made
on many of these, and no precise correlation of the various faunas can be made.
Indeed, the most comprehensive attempt at a post-Tertiary chronology for Australia
is still that of Browne (1945).

While the fragmentary evidence of the upper Tertiary marsupial remains
indicates that the major groups were differentiated in this continent by the Miocene,
many of them reached their acme in the Pleistocene. The fluctuating climate of
that epoch and the consequential rapid changes in the environment no doubt
maintained strong selection pressure, especially on browsing and grazing herbivores

FOSSIL MARSUPIALS AND CAINOZOIC CONTINENTAL STRATIGRAPHY 45

of the open forests and grasslands. The possible results, rapid evolution, with
increasing specialization, gigantism, and extinction, are all evident in the-
palaeontological record of the Quaternary.

The occurrence of any fossil marsupials to the north of the Australian mainland,
is of particular interest, in that information may be gained on the times and directions
of dispersal of the various marsupial groups. To date, their occurrence is restricted
to the diprotodontid, Nototherium watutense , and a few fragmentary undescribed
macropodids from the Morobe Goldfields area. Information kindly supplied by
Dow (1961, pers. commun.) indicates that these fossils are derived from the Otibanda
Lake Beds which were deposited in two lakes in the valleys of the Bulolo and Watut
Rivers, separated by extensive andesitic volcanics of the Bulolo Gorge. The sediments
of the smaller area near Wau comprise conglomerate, sandstone, siltstone, and mud-
stone, with interbedded andesitic tuff and agglomerate near the base. A Pleistocene
age is considered likely for this sequence. The larger lake, in the valleys of the
Watut and lower Bulolo, was formed by faulting and regional uplift in the Snake
River area. The presence of andesitic volcanic material, apparently subaerially
deposited, throughout much of the section, suggests that in part this sequence may
be slightly older. The Otibanda Lake Beds are deformed, with dips up to 45°.

In a small collection recently received from this area the only well preserved
specimen is portion of a diprotodontid mandible referable to Nototherium sp. Its
molar pattern is very similar to Nototherium tasmanicum from the upper Pleistocene^
of Tasmania and the more widely distributed N. mitchelli of the Pleistocene. However,
this comparison does not assist correlation since the isolated upper premolar figured
by Stirton (1957b) suggests that the genus Nototherium ranged at least from upper
Miocene time.

One of the most intensively collected areas of Pleistocene alluvia in Australia
occurs in the valleys of the Condamine River and its tributaries of the eastern
Darling Downs. Many of the type specimens of the Pleistocene marsupials described
by Sir Richard Owen in the last century were obtained from these deposits. In the
eastern part of the area, as in the valley of King Creek, the fossiliferous beds comprise
brown calcareous clays with lenticular basaltic gravels at rather shallow depths in the
creek sections. Along the Condamine River near Dalby and Macalister, the observed
sections are thicker and commonly contain sands and grits as well as brown and grey
clays. Bore records show up to 167 feet of alluvia in this area. A general Pleistocene
age has been assigned to this sequence by Woods (1960) based on the almost complete
absence of living species among the fossils. It is feasible that the more superficial
fossiliferous alluvia along King Creek are upper Pleistocene in age, while the thicker
sediments in the Condamine valley range back to include equivalents of the Chinchilla
Sand.

46

MEMOIRS OF THE QUEENSLAND MUSEUM

Marsupials are predominant in this fauna from the eastern Darling Downs ;
reptiles are not so common as in the Chinchilla Sand. The most common of the giant
marsupials is the widespread Diprotodon optatus, and the dominant marcopodids
are Macropus titan and Protemnodon anak. The smaller polyprotodont marsupials
.and the phalangerids are not well represented, but those with a general forest habitat
and small size are unlikely to be preserved in the fluviatile deposits of wide valleys
and plains.

Following the early work of Owen, a large number of fossil marsupials were
described from the Darling Downs by C. W. De Vis. His descriptions are frequently
unaccompanied by locality and stratigraphic data, and in some cases type specimens
were not designated. This applies, in particular, to his work on the macropodids
(De Vis, 1895), which deals in composite fashion with collections from both the
Chinchilla area and the eastern Darling Downs, and stratigraphic evaluation of many
of his species has not been possible. Currently, revisionary work on these forms is
being undertaken by Dr. W. D. L. Hide and Mr. A. Bartholomai.

Diprotodon optatus was widely distributed on the mainland, and while it
Teached King Island, there is no record of its having reached Tasmania. It was
apparently adapted to a wide range of habitats ; Gill (1955) records its occurrence
at altitudes between sea level and 2,000 feet. The apparent absence of Diprotodon
from Tasmania may be due to a relatively late dispersal of the genus and the
presence of an indigenous species of Notothcrium ( N . tasmanicum) as well as the wide-
spread N. mitchelli in the Mowbray Swamp Peat, regarded by Gill and Banks (1956)
as upper Pleistocene, may be taken as evidence in support of this view. While the
restriction of the range of the family Diprotodontidae in southern Australia to the
upper Pleistocene or the upper part of the middle Pleistocene as suggested by Keble
(1945) is no longer tenable, indications are that most occurrences of Diprotodon
optatus in this region are in upper Pleistocene deposits. Furthermore, the species
is known to range into early Recent time.

However, Diprotodon sp. occurs in the Chinchilla Sand, and Owen (1870)
recorded the genus from a depth of 100 feet in the Condamine alluvia. Its apparent
variation in time range may reflect the local patterns of sedimentation in different
parts of the continent.

Two separate faunas have been recognised by Stirton et al. (1961) from the
Katipiri Sands of Pleistocene age in the Tirari Desert. These fluviatile deposits
rest disconformably on the Tirari Formation. The presumably older mammalian
fauna from Lake Kanunka, tentatively referred by these authors to the early
Pleistocene, contains not only marsupials but the oldest rodent known from the
Australian Region. The marsupials comprise a dasyurid, a thylacoleonid, two
phascolomids including one specimen referable to Phascolonus, several macropodids
■of diverse groups, and diprototodontid fragments, possibly referable to Euowenia.
Diprotodon has not been collected in this fauna.

FOSSIL MARSUPIALS AND CAINOZOIC CONTINENTAL STRATIGRAPHY 47

The other mammalian fauna, placed as late Pleistocene, includes several
rodents, as well as marsupials, comprising the dasyurid Sarcophilus, the phalangerid
Trichosurus, the phascolomid Phascolonus, several macropodids including the still-
living Bettongia lesueuri, and the widespread diprotodontid Diprotodon.

A large assemblage of living and extinct marsupial genera, including Diprotodon,
has been obtained from the lowest unit of the aeolian sequence at the archaeological
site at Lake Menindee (Tedford, 1955). This fauna was contemporaneous with
aboriginal man with the Tartangan culture, and radiocarbon dating of the site
shows it to be of early Recent age at approximately 6,570 years before the present
time (Tindale, 1957). People with the Tartangan culture and those with the earlier
Kartan culture both reached Tasmania, so no zoogeographic basis for the failure of
Diprotodon to reach there is apparent.

Few of the extinct marsupials seemed to have survived the time of the Mid-
Recent Thermal Maximum, about 5,000 years ago. The macropodid Procoptodon
survived until the time of the Pirrian culture of australoid people, about 4,250 years
ago (Tindale, 1957). It would appear that the extinction of Pleistocene marsupial
species was progressive, not a catastrophic result of any sudden climatic change,,
although the demands of the fluctuating enviroment on genetically senile populations
were certainly important factors. The arrival of Man probably had a critical effect
on certain species, especially those of small numbers in restricted areas.

Cave earths and tufas have yielded abundant fossil mammals, including
many not known or poorly represented in other deposits ; but these differences
largely reflect differences in habitat and mode of accumulation of the remains.
Predators, including owls, appear responsible for most of the material. Native rodents
and bats are well represented in some occurrences. The assemblages usually comprise
a mixture of extinct and living species. While many of the deposits are of upper
Pleistocene or even younger age, it is possible that some collections represent material
from more than one stratigraphic horizon. Furthermore, the time ranges of many
living species of small marsupials have not been established, and the taxonomic
evaluation of the fragmentary material often presents difficulties.

Notes on the cave fossils of south-western Western Australia were published
by Glauert (1948) and these embody references to his earlier work in the area.
A radiocarbon date of > 37,000 years for one of these occurrences, the Mammoth
Cave, has been indicated by Ride (1960). In this latter paper the fauna of the
Wombeyan Caves, New South Wales, is discussed and regarded as upper Pleistocene
in age.

Studies of cave faunas of Recent age in southern Western Australia by
Lundelius (1957) show that the geographic ranges of many living species were vastly
different in prehistoric time. Very recent material from caves in the Buchan district

48

MEMOIRS OF THE QUEENSLAND MUSEUM

■of Victoria, listed by Wakefield (1960), is of particular interest in that it includes
Burramys parvus, previously known only from an assemblage of living and extinct
species from the Wombeyan Caves.

As biostratigraphic studies involving our Quaternary mammals proceed, it will
be necessary to give special consideration to the local time ranges of species, which
vary with time of dispersal and time of extinction. This variation becomes of greater
significance with the decrease in age of the fauna.

Mr. Duncan Dow, of the Bureau of Mineral Resources, kindly furnished
information on the stratigraphy of the Otibanda Lake Beds, in the Morobe Goldfields
-area, Territory of New Guinea.

LITERATURE CITED

Anderson, C., 1937. Palaeontological Notes. No. IV. Bee. Aust. Mus., 20, pp. 73-78.

Browne, W. R., 1945. An Attempted Post-Tertiary Chronology for Australia. Proc. Linn. Soc.
N.S. W., 70, pp. v-xxiv.

J)e Vis, C. W., 1895. A Review of the fossil jaws of the Macropodidae in the Queensland Museum.
Proc. Linn. Soc. N.S.W., 10, pp. 75-133.

Dun, W. S., 1895. Notes on the Occurrence of Monotreme Remains in the Pliocene of New
South Wales. Rec. Geol. Surv. N.S.W., 4, pp. 118-126.

•Gill, E. D., 1955. The Problem of Extinction with Special Reference to the Australian
Marsupials. Evolution, 9, pp. 87-92.

, 1957. The Stratigraphical Occurrence and Palaeoecology of some Australian

Tertiary Marsupials. Mem. Nat. Mus. Vic., 21, pp. 135-199.

, and Banks, M. R., 1956. Cainozoic History of Mowbray Swamp and other Areas

of North-western Tasmania. Rec. Queen Viet. Mus. Launc., n.s. 6, pp. 1-41.

Glaessner, M. F., and McGowran, B., and Wade, M., 1960. Discovery of a Kangaroo Bone in the
Middle Miocene of Victoria. Aust. J. Sci., 22, pp. 484-485.

Glauert, L., 1948. The Cave Fossils of the South-West. W. Aust. Nat., 1, pp. 101-104.

Keble, R. A., 1945. The Stratigraphic Range and Habitat of the Diprotodontidae in Southern
Australia. Proc. Roy. Soc. Viet., 57, pp. 23-48.

King, L., 1950. The Cyclic Land-surfaces of Australia. Proc. Roy. Soc. Viet., 62, pp. 79-95.

Lundelius, E., 1957. Additions to Knowledge of the Ranges of Western Australian Mammals.
W. Aust. Nat., 5, pp. 173-182.

Gwen, R., 1870. On the Fossil Mammals of Australia. Part III. Phil. Trans., 160, pp. 519-578.

FOSSIL MARSUPIALS AND CAINOZOIC CONTINENTAL STRATIGRAPHY 49

Paten, R. J., 1961. The Tertiary Geology of Western Queensland. Bur. Min. Resour. Aust.
Rec., 1961/52, pp. 1-28.

Ride, W. D. L., 1960. The Fossil Mammalian Fauna of the Burramys parvus Breccia from the
Wombeyan Caves, New South Wales. J. Roy. Soc. W. Aust., 43, p. 74.

Stirton, R. A., 1955. Late Tertiary Marsupials from South Australia. Rec. S. Aust. Mus., 11,
pp. 247-268.

, 1957a. A new koala from the Pliocene Palankarinna fauna of South Australia.

Rec. S. Aust. Mus., 13, pp. 73-81.

, 1957b. Tertiary Marsupials from Victoria, Australia. Mem. Nat. Mus. Viet., 21,

pp. 121-134.

, Tedford, R. H., and Miller, A. H., 1961. Cenozoic Stratigraphy and Vertebrate

Paleontology of the Tirari Desert, South Australia. Rec. S. Aust. Mus., 14, pp. 19-61.

Tedford, R. H., 1955. Report on the Extinct Mammalian Remains at Lake Menindee, New South
Wales. Rec. S. Aust. Mus., 11, pp. 200-305.

Tindale, N. B., 1957. Culture Succession in South-Eastern Australia from Late Pleistocene to
the Present. Rec. S. Aust. Mus., 13, pp. 1-50.

Twidale, C. R., 1956. Chronology of Denudation in North-West Queensland. Bull. Geol. Soc.
Amer., 67, pp. 867-882.

Wakefield, N., 1960. Recent Mammal Bones from the Buchan District — 1. Viet. Nat., 77,
pp. 164-178.

Wood Jones, F., 1931. A Re-examination of the Skeletal Characters of Wynyardia bassiana, an
extinct Tasmanian Marsupial. Pap. dk Proc. Roy. Soc. Tas. for 1930, pp. 99-115.

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S. G. Reid, Government Printer, Brisbane

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